Kevin A. Wood, Phoebe Ham, Jake Scales, Eleanor Wyeth, Paul E. Rose. 2020: Aggressive behavioural interactions between swans (Cygnus spp.) and other waterbirds during winter: a webcam-based study. Avian Research, 11(1): 30. DOI: 10.1186/s40657-020-00216-7
Citation: Kevin A. Wood, Phoebe Ham, Jake Scales, Eleanor Wyeth, Paul E. Rose. 2020: Aggressive behavioural interactions between swans (Cygnus spp.) and other waterbirds during winter: a webcam-based study. Avian Research, 11(1): 30. DOI: 10.1186/s40657-020-00216-7

Aggressive behavioural interactions between swans (Cygnus spp.) and other waterbirds during winter: a webcam-based study

More Information
  • Corresponding author:

    Paul E. Rose, p.rose@exeter.ac.uk

  • Received Date: 28 Jun 2020
  • Accepted Date: 01 Aug 2020
  • Available Online: 24 Apr 2022
  • Publish Date: 06 Aug 2020
  • Background 

    Our understanding of any impacts of swans on other waterbirds (including other swans), and potential effects on waterbird community structure, remain limited by a paucity of fundamental behavioural and ecological data, including which species swans interact aggressively with and how frequently such interactions occur.

    Methods 

    Behavioural observations of aggression by swans and other waterbirds in winters 2018/2019 and 2019/2020, were carried out via live-streaming webcams at two wintering sites in the UK. All occurrence sampling was used to identify all aggressive interactions between conspecific or heterospecifics individuals, whilst focal observations were used to record the total time spent by swans on aggressive interactions with other swans. Binomial tests were then used to assess whether the proportion of intraspecific aggressive interactions of each species differed from 0.5 (which would indicate equal numbers of intraspecific and interspecific interactions). Zero-inflated generalized linear mixed effects models (ZIGLMMs) were used to assess between-individual variation in the total time spent by swans on aggressive interactions with other swans.

    Results 

    All three swan species were most frequently aggressive towards, and received most aggression from, their conspecifics. Our 10-min focal observations showed that Whooper (Cygnus cygnus) and Bewick's Swans (C. columbianus bewickii) spent 13.8± 4.7 s (means± 95% CI) and 1.4± 0.3 s, respectively, on aggression with other swans. These durations were equivalent to 2.3% and 0.2% of the Whooper and Bewick's Swan time-activity budgets, respectively. Model selection indicated that the time spent in aggressive interactions with other swans was best-explained by the number of other swans present for Whooper Swans, and an interactive effect of time of day and winter of observation for Bewick's Swans. However, the relationship between swan numbers and Whooper Swan aggression times was not strong (R2= 19.3%).

    Conclusions 

    Whilst swans do exhibit some aggression towards smaller waterbirds, the majority of aggression by swans is directed towards other swans. Aggression focused on conspecifics likely reflects greater overlap in resource use, and hence higher potential for competition, between individuals of the same species. Our study provides an example of how questions relating to avian behaviour can be addressed using methods of remote data collection such as live-streaming webcams.

  • In January (K.L. Chang) and May 2010 (J.Y. Lei and L. Chen), we carried out two separate birding trips in the southern Zhangmu, a small town in southern Tibet (27°59′37′′N, 85°58′00′′E; 1755–2380 m a.s.l.) adjacent to Nepal. We observed birds by binoculars, made photographs and recorded sounds of some significant bird species during these trips. Among the bird species we recorded in these trips, we identified the Himalayan Black-lored Tit (Parus xanthogenys) and the Nepal Wren-Babbler (Pnoepyga immaculate) which have not been included in any checklist of birds of China (e.g. Cheng, 1987; MacKinnon et al., 2000; Zheng, 2005) and hence they are confirmed as new bird records for China. Here we describe details of these two new bird records.

    Two tits with vivid yellow facial patterns were recorded by K.L. Chang on 18 January 2010 and were quickly identified as Himalayan Black-lored Tits according to his previous birding experience of this species in Nepal (Fig. 1). In a subsequent spring trip, J.Y. Lei, L. Chen and four other observers also located and photographed a flock of Himalayan Black-lored Tits (n > 10) on the road from Zhangmu to the China-Nepal border checkpoint at an elevation of 1600 m on 12 May and 15 May 2010. This species may be either a common or an uncommon resident at Zhangmu.

    Figure  1.  The Himalayan Black-lored Tit (Parus xanthogenys) (Photo by Liang Chen)

    The Himalayan Black-lored Tit (Fig. 1) is a large-sized tit, estimated to be about 13 cm in length, with a body size equally large as that of the Great Tit (Parus major) and the Yellow-cheeked Tit (P. spilonotus). It is a yellow-and-black tit, with a broad black line through the middle of its underparts, extending to the breast and throat. The long and pronounced crest, neck and front are also black with yellow cheeks. Black stripes, started from the neck, extend to the lores, constituting a striking facial pattern. The upper parts are olive-green with a few black streaks confined to the scapulars. It resembles the Yellow-cheeked Tit (P. spilonotus), probably its closest relative, as well as the other yellow-and-crested tit in the Himalayas, but this latter bird apparently lacks black in its forehead and lores. Females and young birds are reported to be duller in color than males (Grimmett et al., 1999; Rasmussen and Anderton, 2005). The flock we observed may have been a mixture of both males and female-like birds.

    This species is a common resident in open tropical forests, secondary forests and edges of dense forests and groves of the Indian subcontinent, from north Pakistan through the Himalayas to eastern Nepal (Grimmett et al., 1999; Rasmussen and Anderton, 2005). Three subspecies were described under the species level, the nominate races xanthogenys, aplonotus and travancoreensis (Grimmett et al., 1999; Rasmussen and Anderton, 2005). Based on evidence of morphology and vocalization, Rasmussen and Anderton (2005) proposed that xanthogenys and the two other subspecies should be treated as two species: the Himalayan Black-lored Tit (Parus xanthogenys) (monotypic) and the India Black-lored Tit (P. aplonotus), with subspecies aplonotus and travancoreensis. Gill (2010) adopted this taxonomic treatment in the IOC World Bird Names. The location where we sighted the birds, our field description and morphological characteristics shown in the photographs, explicitly match the diagnosis of the taxon xanthogenys. Following Gill (2010), we therefore confirm that our sighting of the Himalayan Black-lored Tit (P. xanthogenys) is a new record for this bird species in China.

    On 8 May 2010, near the Zhangmu Hotel we heard a series of high-pitched metallic notes, like "tsi-tsu-tsi-tsi-tsu-tsi-tsi-tsi", speeding up at the end and repeating several times. We then recorded its repeating sound and did play-back. The bird responded promptly and was easily lured out, standing at a fallen twig and presenting a full view to us, and was identified as a Nepal Wren-Babbler. It was a typical small, round and tailless Wren-Babbler, about 10 cm long. The bird was uniformly fulvous on its upper and under parts with dense black arrow-shaped streaks on breast and belly. Moreover, the bird did not have eye-brows.

    There are three potential wren-babbler species of such a morphological type, sympatrically occurring in the Himalayas: the Scaly-breasted Wren-Babbler (Pnoepyga albiventer), the Pygmy Wren-Babbler (P. pusilla) and the Nepal Wren-Babbler (P. immaculate). Considering the species we observed at Zhangmu, we first ruled out P. pusilla by its unspotted wings and tertials (Fig. 2). We are also very familiar with the diagnostic "three-note" song by P. pusilla, which we did not hear in this case. With respect to P. albiventer, our species apparently lacks spots on its crown, mantle and wing-covers (Fig. 2). We are therefore inclined to infer that the bird is a Nepal Wren-Babbler (P. immaculate). This species is reported to have a longer and heavier bill than P. pusilla (Grimmett et al., 1999; Rasmussen and Anderton, 2005), but it was almost impossible for us, on the basis of this trait, to make detailed comparisons in these field conditions. Afterwards, we compared our derived song type to a reference song of a putative Nepal Wren-Babbler (available at http://www.xeno-canto.org/browse.php?query=Nepal+Babbler) and consolidated our preliminary judgment about the species identification. During the next few days (until 18 May), we repeatedly heard this distinctive song at many other sites between 1800–2200 m elevation around Zhangmu County, indicating that the species might be a fairly common breeder in the region.

    Figure  2.  The Nepal Wren-Babbler (Pnoepyga immaculate) (Photo by Haitao Li)

    The Nepal Wren-Babbler has been neglected as a full species because its song was long thought to be a variant of the song of the Scaly-breasted Wren-Babbler (P. albiventer) (Martens and Eck, 1991). It has lately been realized that birds with this kind of song are morphologically differentiated from the Scaly-breasted Wren-Babbler (unspotted crown, mantle and wing-covers) and hence, the status of this unknown bird species was resolved. The Nepal Wren-Babbler is an endemic resident breeder of the Indian Subcontinent and the Himalayas, from eastern Nepal west to Himachal Pradesh in India. It is commonly found in temperate forests and subtropical or tropical moist montane habitats in undergrowth near streams and rivers (Grimmett et al., 1999; Rasmussen and Anderton, 2005). There are no records of this species in Tibet, China in any published checklist of birds of China, which is presumably due to wrong species identification resulting from neglect of subtle differences between this species and its close relatives.

    Although Zhangmu is a small town, it has become a hot birding spot in China since it has already witnessed several new records of bird species for China in recent years, including the Himalayan Bulbuls (Pycnonotus leucogenys) (Zhang et al., 2006), Tickell's Thrush (Turdus unicolor) (Yu, 2008) and the Black-chinned Babbler (Stachyridopsis pyrrhops) (Dong and Yang, 2010). More new bird species for China are expected to be recorded at this site in the future.

    We thank Yang Liu and Yat-Tung Yu for their constructive comments, which substantially improved an earlier draft. We are grateful to two anonymous reviewers who provided valuable comments on an earlier draft of this manuscript. We also thank Haitao Li, Qin Wang, Li Zhang and Jia Zhong for joining our field expeditions.

  • Altmann J. Observational study of behavior: sampling methods. Behaviour. 1974;49:227–66.
    Amat JA. Food usurpation by waterfowl and waders. Wildfowl. 1990;41:107–16.
    Anderson MJ, Urbine JL, Wilson C, Callabro L. Employment of web-based images and a live web cam in the examination of lateral neck-resting preferences in the American flamingo (Phoenicopterus ruber). J Caribb Ornithol. 2011;24:41–7.
    Arnold TW. Uninformative parameters and model selection using Akaike's Information Criterion. J Wildl Manage. 2010;74:1175–8.
    Bailey RO, Batt BDJ. Hierarchy of waterfowl feeding with Whistling Swans. Auk. 1974;91:488–93.
    Barton K. MuMIn: Multi-Model Inference. Version 1.43.15. 2019. .
    Beekman J, Koffijberg K, Wahl J, Kowallik C, Hall C, Devos K, et al. Long-term population trends and shifts in distribution for Bewick's Swans Cygnus columbianus bewickii wintering in northwest Europe. Wildfowl. 2019; Special Issue 5: 73–102.
    Beven G. Coot feeding on weed disturbed by Mute Swans. Brit Birds. 1980;73:219–20.
    Black JM, Rees EC. The structure and behaviour of the Whooper Swan population wintering at Caerlaverock, Dumfries and Galloway, Scotland: an introductory study. Wildfowl. 1984;35:21–36.
    Bowler JM. Feeding strategies of Bewick's Swans (Cygnus columbianus bewickii) in winter. PhD Thesis. Bristol: University of Bristol; 1996.
    Brazil MA. A case of unusual aggression by a Whooper Swan. Tori. 1983;32:155.
    Brides K, Wood KA, Hearn RD, Fijen TPM. Changes in the sex ratio of the Common Pochard Aythya ferina in Europe and North Africa. Wildfowl. 2017;67:100–12.
    Brooks ME, Kristensen K, van Benthem KJ, Magnusson A, Berg CW, Nielsen A, et al. glmmTMB balances speed and flexibility among packages for zeroinflated generalized linear mixed modeling. R J. 2017;9:378–400.
    Burgess RM, Stickney AA. Interspecific aggression by Tundra Swans towards Snow Geese on the Sagavanirktok River Delta, Alaska. Auk. 1994;111:204–7.
    Burnham KP, Anderson DR, Huyvaert KP. AIC model selection and multimodel inference in behavioral ecology: some background, observations, and comparisons. Behav Ecol Sociobiol. 2011;65:23–35.
    Clopper CJ, Pearson ES. The use of confidence or fiducial limits illustrated in the case of the binomial. Biometrika. 1934;26:404–13.
    Conover MR, Kania GS. Impact of interspecific aggression and herbivory by mute swans on native waterfowl and aquatic vegetation in New England. Auk. 1994;111:744–8.
    Crawley MJ. The R Book. 2nd ed Chichester: Wiley; 2013.
    Davis JB, Guillemain M, Kaminski RM, Arzel C, Eadie JM, Rees EC. Habitat and resource use by waterfowl in the northern hemisphere in autumn and winter. Wildfowl. 2014;4:17–69.
    Delacour J. Waterfowl in large mixed collections. Int Zoo Yearb. 1973;13:15–9.
    Dormann CF, Elith J, Bacher S, Buchmann C, Carl G, Carré G, et al. Collinearity: a review of methods to deal with it and a simulation study evaluating their performance. Ecography. 2013;36:27–46.
    Eichorst B. Internet webcams provide opportunities for college student research on animal behavior and ecology: an example with birds. Am Biol Teacher. 2018;80:680–5.
    Ely CR, Budeau DA, Swain UG. Aggressive encounters between Tundra Swans and Greater White-fronted Geese during brood rearing. Condor. 1987;89:420–2.
    Frost TM, Calbrade NA, Birtles GA, Mellan HJ, Hall C, Robinson AE, et al. Waterbirds in the UK 2018/2019: The Wetland Bird Survey. Thetford: BTO/RSPB/ JNCC; 2020.
    Gayet G, Guillemain M, Mesléard F, Fritz H, Vaux V, Broyer J. Are Mute Swans (Cygnus olor) really limiting fishpond use by waterbirds in the Dombes, Eastern France? J Ornithol. 2011;152:45–53.
    Gayet G, Calenge C, Broyer J, Mesléard F, Vaux V, Fritz H, et al. Analysis of spatial point pattern shows no desertion of breeding Mute Swan areas by the other waterbirds within fishpond. Acta Ornithol. 2016;5:151–63.
    Gillham ME. Feeding habits and seasonal movements of mute swans on two south Devon estuaries. Bird Study. 1956;3:205–12.
    Gurtovaya EN. Aggressive interactions between Bewick's Swans and other Anseriformes in the breeding period. Casarca. 2000;6:167–76.
    Gyimesi A, Stillman RA, Nolet BA. Cryptic interference competition in swans foraging on cryptic prey. Anim Behav. 2010;80:791–7.
    Gyimesi A, van Lith B, Nolet BA. Commensal foraging with Bewick's Swans Cygnus bewickii doubles instantaneous intake rate of Common Pochard Aythya ferina. Ardea. 2012;100:55–62.
    Holm S. A simple sequentially rejective multiple test procedure. Scand J Stat. 1979;6:65–70.
    Johnsgard PA. Handbook of waterfowl behavior. New York: Cornell University Press; 1965.
    Källander H. Commensal association of waterfowl with feeding swans. Waterbirds. 2005;28:326–31.
    King JA. The ecology of aggressive behavior. Annu Rev Ecol Syst. 1973;4:117–38.
    Lenth R. Emmeans: Estimated Marginal Means, aka Least-Squares Means. Version 1.4.5. 2020. .
    Lind H. The rotation display of the Mute Swan Cygnus olor: synchronised neighbour responses as instrument in the territorial defence strategy. Ornis Scand. 1984;15:98–104.
    Lüdecke D. Sjstats: Statistical Functions for Regression Models. Version 0.17.9. 2020. .
    Lumsden HG. Trumpeter Swans and Mute Swans compete for space in Ontario. Ontario Birds. 2016;34:14–23.
    Mac Nally R, Duncan RP, Thomson JR, Yen JD. Model selection using information criteria, but is the "best" model any good? J Appl Ecol. 2018;55:1441–4.
    Marchowski D, Neubauer G. Kleptoparasitic strategies of Mallards towards conspecifics and Eurasian Coots. Ardea. 2019;107:110–4.
    Metcalfe NB, Furness RW. Aggression in shorebirds in relation to flock density and composition. Ibis. 1987;129:553–63.
    Nakagawa S, Johnson PC, Schielzeth H. The coefficient of determination R2 and intra-class correlation coefficient from generalized linear mixedeffects models revisited and expanded. J R Soc Int. 2017;14:20170213.
    Newth JL, McDonald RA, Wood KA, Rees EC, Semenov I, Chistyakov A, et al. Predicting intention to hunt protected wildlife: a case study of Bewick's swans in the European Russian Arctic. Oryx. (in press).
    O'Hare MT, Stillman RA, McDonnell JO, Wood LR. Effects of mute swan grazing on a keystone macrophyte. Freshwater Biol. 2007;52:2463–75.
    Peiman K, Robinson B. Ecology and evolution of resource-related heterospecific aggression. Quart Rev Biol. 2010;85:133–58.
    Pelligrini AD. The roles of aggressive and affiliative behaviors in resource control: a behavioral ecological perspective. Dev Rev. 2008;28:461–87.
    Peluso AI, Royer EA, Wall MJ, Anderson MJ. The relationship between environmental factors and flamingo aggression examined with internet resources. Avian Biol Res. 2013;6:215–20.
    Pöysä H. Resource utilization pattern and guild structure in a waterfowl community. Oikos. 1983;40:295–307.
    R Core Team. R: A language and environment for statistical computing. [3.6.3]. Vienna, Austria: R Foundation for Statistical Computing; 2020.
    Rees EC. Bewick's Swan. London: T & AD Poyser; 2006.
    Rees EC, Cao L, Clausen P, Coleman JT, Cornely J, Einarsson O, et al. Conservation status of the world's swan populations, Cygnus sp. and Coscoroba sp.: a review of current trends and gaps in knowledge. Wildfowl. 2019; Special Issue 5: 35–72.
    Richards SA. Dealing with overdispersed count data in applied ecology. J Appl Ecol. 2008;45:218–27.
    RSPB. Birds A- Z: Bird Guides. Royal Society for the Protection of Birds. . Accessed October 2018.
    Schulwitz SE, Spurling DP, Davis TS, McClure CJW. Webcams as an untapped opportunity to conduct citizen science: siz years of the American Kestrel Partnership's KestrelCam. Glob Ecol Conserv. 2018;15:e00434.
    Scott DK. Social behaviour of wintering Cygnus columbianus bewickii. In: Matthews GVT, Smart M, editors. Proceedings of the Second International IWRB Swan Symposium, Sapporo, Japan, 1980. Slimbridge: International Waterfowl Research Bureau; 1981. p. 211–25.
    Shimada T. Ducks foraging on swan faeces. Wildfowl. 2012;62:224–7.
    Sladen WJL. Swans should not be hunted. In: Sears J, Bacon PJ, editors. Proceedings of the Third International IWRB Swan Symposium, Oxford, 1989. Slimbridge: International Waterfowl Research Bureau; 1991. p. 368–75.
    Stone WB, Marsters AD. Aggression among captive Mute Swans. New York Fish Game J. 1970;17:51–3.
    Tatu KS, Anderson JT, Hindman LJ, Seidel G. Diurnal foraging activities of mute swans in Chesapeake Bay, Maryland. Waterbirds. 2007;30:121–9.
    Therres GD, Brinkler DF. Mute Swan interactions with other birds in Chesapeake Bay. In: Perry MC, editor. Mute Swans and Their Chesapeake Bay Habitats: Proceedings of a Symposium. Virginia: US Geological Survey; 2004. p. 43–6.
    Tingay A. Aggression in the Black Swan. Emu. 1974;74:35–8.
    Vogrin M. A Coot Fulica atra eating waterfowl droppings. Butll GCA. 1997;14:63–4.
    Włodarczyk R, Minias P. Division of parental duties confirms a need for biparental care in a precocial bird, the mute swan Cygnus olor. Anim Biol. 2015;65:163–76.
    Wood KA, Stillman RA, Goss-Custard JD. The effect of kleptoparasite and host numbers on the risk of food-stealing in an avian assemblage. J Avian Biol. 2015;46:589–96.
    Wood KA, Ponting J, D'Costa N, Newth JL, Rose PE, Glazov P, et al. Understanding intrinsic and extrinsic drivers of aggressive behaviour in waterbird assemblages: a meta-analysis. Anim Behav. 2017;126:209–16.
    Wood KA, Cao L, Clausen P, Ely CR, Luigujõe L, Rees EC, et al. Current trends and future directions in swan research: insights from the 6th International Swan Symposium. Wildfowl. 2019a; Special Issue 5:1–34.
    Wood KA, Hilton GM, Newth JL, Rees EC. Seasonal variation in energy gain explains patterns of resource use by avian herbivores in an agricultural landscape: insights from a mechanistic model. Ecol Model. 2019b;409:108762.
    Zuur AF, Ieno EN, Elphick CS. A protocol for data exploration to avoid common statistical problems. Methods Ecol Evol. 2010;1:3–14.
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